2011 gene list for melon - cucurbit breedingcuke.hort.ncsu.edu/cgc/cgc3334/cgc3334-24.pdf · 2011...

104 / Cucurbit Genetics Cooperative Report 33-34: 104-133 (2010-2011)

2011 Gene List for MelonCatherine DogimontINRA, UR1052, Unité de Génétique et d’Amélioration des Fruits et Légumes, BP 94, 84143 Montfavet cedex (France)[email protected]

Melon (Cucumis melo L.) is an economically impor-tant, cross-pollinated species. Melon has 2n = 24 chro-mosomes and a relatively small genome (450 Mb), aboutthree times larger than the Arabidopsis thaliana genomeand similar to the rice genome (Arumanagathan andEarle, 1991). Melon has high intra-specific genetic varia-tion and morphologic diversity. A great variety of ge-netic and molecular studies have been conducted onimportant agronomical traits, such as resistance topathogens and insects, and floral and fruit traits.

The following list is the latest version of the genelist for melon. Previous gene lists were organized byPitrat: (Pitrat, 2006), (Pitrat, 2002), (Pitrat, 1998), (Pitrat,1994), (Pitrat, 1990), (Pitrat, 1986), (Committee, 1982).This current list has been modified from previous listsin that (1) it provides an update of the known genes andQTLs, and (2) it adds an expanded description for re-ported genes including sources of resistance and resis-tance genes, phenotypes of mutants and traits related toseeds, seedlings, plant morphology and architecture,flowers and fruits. Locations of the reported genes onthe melon genetic map and linked markers useful formarker assisted selection were reported where available.

Since the first molecular marker-based melon mappublished in 1996 (Baudracco-Arnas and Pitrat, 1996),several genetic maps of melon have been published byseveral research teams, using several segregating popu-lations. 2011 will be the year of the publication of anintegrated map of melon in the framework of the Inter-national Cucurbit Genomic Initiative (Díaz et al., 2011).The integrated map has been constructed by mergingdata from eight independent mapping populations us-ing genetically diverse parental lines. It spans 1150 cMdistributed across the 12 melon linkage groups and com-prises more than 1500 markers. Individual maps andthe integrated map are available at www.icugi.org. Thelinkage groups were named according to Perin et al.,2002. The same nomenclature will be adopted hereafter.QTLs for 62 traits including virus resistance, fruit shape,fruit weight, sugar content have been located on thisintegrated map (Díaz et al., 2011).

The list of melon sequences was not updated, asmelon ESTs and full cDNAs are increasing extraordi-narily (www.icugi.org). A physical map of the melongenome, anchored to the genetic map, has been estab-

lished (Gonzalez et al., 2010) and the complete melongenome sequence is expected by the end of the year.

Host Plant Resistance genesConsiderable attention has been given to resistance

genes in melon. Genes for resistance to viruses, insects,fungi and oomycetes have been reported.

Viral DiseasesThe first source for resistance to Zucchini yellow

mosaic virus (ZYMV, Potyvirus), and for a long time theonly known source, was the Indian accession PI 414723(Pitrat et al., 1996). The resistance proved to be strain-specific and was not effective against a second pathotypeof the virus. The screening of about 60 cultivars fromIran allowed the identification of three immune culti-vars: Magolalena Vertbrod, Soski and Bahramabadi(Arzani and Ahoonmanesh, 2000). Among 200 melonscollected in Sudan, resistance sources to ZYMV werefound, mainly in wild forms (Mohamed, 1999).

Resistance to ZYMV in PI 414723 was reported tobe controlled by a single dominant gene, Zym (Pitrat andLecoq, 1984), which mapped to the linkage group II(former LG 4), linked to the gene a (andromonoecious)(Pitrat, 1991; Perin et al., 2002). Using the ZYMV-Natstrain (pathotype 1), Danin-Poleg et al. (1997) found thatthree genes were needed to confer the resistance in PI414723 (Zym-1, Zym-2 and Zym-3). Molecular markerslinked to the resistance were identified by bulk segregantanalysis (Danin-Poleg et al., 2000; Danin-Poleg et al.,2002).

A semi-dominant gene named Fn, independent ofZym, was reported to control in ‘Doublon’ plant wiltingand necrosis after inoculation with strains of the Fpathotype of ZYMV (Risser et al., 1981). The Fn genewas located in the linkage group V (formerly 2), at 12 cMof the Vat gene, conferring Aphis gossypii resistance(Pitrat, 1991).

Necrosis after inoculation with Watermelon mosaicvirus-Morocco (Potyvirus) was reported to be controlledby a single dominant gene Nm in ‘Védrantais’ (nm in‘Ouzbèque’) (Quiot-Douine et al., 1988).

Papaya ringspot virus- watermelon type (PRSV, for-merly called WMV-1, Potyvirus) resistance was reported

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in the Indian accessions PI 180280 (Webb and Bohn,1962; Webb, 1979), PI 180283 (Quiot et al., 1971), PI414723 (Anagnostou and Kyle, 1996), and PI 124112(McCreight and Fashing-Burdette, 1996) and in TGR-1551 (C-105) from Zimbabwe (Gómez-Guillamón et al.,1998). Resistance to PRSV-W is conferred by a singledominant gene, Prv, in PI 180280 (Webb, 1979) as wellas in the lines B66-5 and WMR 29, derived from PI180280 (Pitrat and Lecoq, 1983). An allele at the samelocus was shown to incite a lethal necrotic responseagainst French strains of PRSV-W in PI 180283 and in72025, derived from PI 180283 (Pitrat and Lecoq, 1983).These alleles were called Prv1 and Prv2, Prv1 being domi-nant over Prv2 (Pitrat, 1986). Prv has been mapped to thelinkage group IX (former 5) (Pitrat, 1991; Perin et al.,2002), closely linked to the gene Fom-1 conferring resis-tance to Fusarium oxysporum races 0 and 2 (Pitrat, 1991;Perin et al., 2002; Brotman et al., 2005). A single domi-nant gene, Prv-2, was also reported to control an incom-patible reaction of PI 124112 after inoculation with PRSV(McCreight and Fashing-Burdette, 1996).

Partial resistance to Watermelon mosaic virus (WMV,formerly WMV-2, Potyvirus) has been reported in melonline 91213, which was selected from PI 371795 and re-lated to PI 414723 (Moyer et al., 1985; Gray et al., 1988;Moyer, 1989), the Korean accession PI 161375 (Pitrat,1978), and in the accessions from Iran (Latifah-1,Tashkandi and Khorasgani) and in an exotic line(Galicum) (Arzani and Ahoonmanesh, 2000). Partialresistance was reported in breeding lines obtained bysuccessive backcrossing with selection from PI 414723;inoculated plants develop mosaic symptoms on inocu-lated leaves but recover from symptoms and virus infec-tion in the youngest leaves. This partial resistance wasreported to be controlled by a single dominant gene, Wmr,linked to the ZYMV resistance gene, Zym (Gilbert et al.,1994; Anagnostou et al., 2000). PI 414723 was observedto be highly susceptible to WMV after inoculation withEuropean strains of WMV (Dogimont et al., unpublisheddata; Gómez-Guillamón, 1998). The accession TGR-1551was reported to exhibit very mild symptoms and a veryreduced titer of virus; this partial resistance was essen-tially determined by a recessive gene (Diaz-Pendon etal., 2005). Still unnamed, we propose to name it wmr-2.

Several cultivars originating from Asia and belong-ing to Oriental pickling melon (var. conomon) and to Ori-ental melon (var. makuwa) were reported to be highly re-sistant to Cucumber mosaic virus (CMV, Cucumovirus)(Enzie, 1943; Webb and Bohn, 1962; Risser et al., 1977;Hirai and Amemiya, 1989; Daryono et al., 2003; Diaz etal., 2003). Interestingly, some accessions from Iran werealso found resistant to CMV (Arzani and Ahoonmanesh,2000) as well as the Indian IC274014 (Dhillon et al., 2007).

Resistance to the CMV-B2 strain in the accessionYamatouri was reported to be controlled in a single domi-nant manner. SCAR markers linked to the gene, namedCreb-2, were identified (Daryono et al., 2010). CMV resis-tance was first reported to be controlled by three reces-sive genes in the cross Freeman Cucumber x Noy Amid(Karchi et al., 1975). Seven QTLs were shown to be in-volved in resistance to three different strains of CMV inthe cross Védrantais x PI 161375 (Dogimont et al., 2000);one of them, located in linkage group XII explains a largepart of the resistance to the strain P9 (Dogimont et al.,2000; Essafi et al., 2009).

Among about 500 accessions tested, resistance toCucurbit aphid borne yellows virus (CABYV, Polerovirus,transmitted by aphids on a persistent manner) was re-ported in the Indian accessions 90625 (= PI 313970),Faizabadi Phoont, PI 124112, PI 282448, and PI 414723,in the Korean accession PI 255478 and in PI 124440 fromSouth Africa (Dogimont et al., 1996). Resistance toCABYV in PI 124112 is conferred by two independentcomplementary recessive genes, named cab-1 and cab-2(Dogimont et al., 1997).

Partial resistance to the Beet pseudo yellows virus(BPYV, Crinivirus), transmitted by the whiteflyTrialeurodes vaporariorum, was reported in a few acces-sions of Asian origin: Nagata Kim Makuwa, PI 161375,Cma, a wild melon collected in Northern Korea and aSpanish landrace Tendral type (Esteva et al., 1989; Nuezet al., 1991). The resistance of Cma, expressed as a de-layed and milder infection, resulted from the cumula-tive effect of an antixenosis against the vector and resis-tance to the virus (Soria et al., 1996; Nuez et al., 1999).Study of segregating families under natural infectionsuggested that the partial resistance to BPYV in NagataKim Makuwa, PI 161375 and Cma was controlled bysingle genes, partially dominant in Nagata Kim Makuwa(gene My) and Cma, and partially recessive in PI 161375(Esteva and Nuez, 1992; Nuez et al., 1999).

Resistance to Cucurbit yellow stunting disorder virus(CYSDV, Crinivirus) was reported in the accession TGR-1551 (C-105), from Zimbabwe, under natural infectionin Spain and when subjected to controlled inoculationby viruliferous Bemisia tabaci and by grafting (Lopez-Seseand Gomez-Guillamon, 2000). Delayed and only slightsymptoms were reported in a few accessions under natu-ral infection conditions in the United Arab Emirates (Ju-piter, Muskotaly, PI 403994) and in Spain (Hassan et al.,1991; Lopez-Sese and Gomez-Guillamon, 2000). Partialresistance to CYSDV was also reported in PI 313970 inthe United States (McCreight and Wintermantel, 2008).In progenies obtained from the cross between TGR-1551and a susceptible Spanish Piel de Sapo cultivar, the re-sistance was shown to be controlled by a single domi-


nant gene, called Cys (Lopez-Sese and Gomez-Guillamon, 2000).

A large melon germplasm was tested for Lettuceinfectious yellows virus (LIYV, Crinivirus) resistance innatural infection by Bemisia tabaci biotype A in Califor-nia. A snake melon originating from Saudi Arabia wasshown to exhibit very mild LIYV symptoms (McCreight,1991; McCreight, 1992). After successive field tests andconfirmation in controlled-inoculation greenhouse tests,the Indian accession PI 313970, was shown to be themost interesting source of resistance to LIYV, althoughan occasional plant of this accession may appear symp-tomatic, or have a positive ELISA for LIYV (McCreight,1998, 2000). Resistance to LIYV in PI 313970 was showncontrolled by a single dominant allele at the locus desig-nated Liy (McCreight, 2000).

Melon breeding line MR-1 and PI 124112, PI179901, PI 234607, PI 313970 and PI 414723 were re-ported to exhibit a partial resistance to Cucurbit leafcrumple virus (CuLCrV), a geminivirus transmitted by B.tabaci biotype B, while PI 236355 was found to be com-pletely resistant. A single recessive gene, named culcrv,was reported to control resistance in PI 313970, and likelyin the other resistant accessions (McCreight et al., 2008).

Gonzalez-Garza et al. reported three phenotypeswhen they inoculated various melon cultivars with Melonnecrotic spot virus (MNSV, Carmovirus) (Gonzalez-Garzaet al., 1979): - cultivars susceptible to systemic infectionshowing local lesions on the inoculated leaves followedby systemic necrotic spotting, necrotic streaks on stems,conducting finally infected plants to collapse; - culti-vars showing local lesions but no systemic symptoms:53% of the accessions tested; - immune lines remainingfree of symptoms (‘Improved Gulfstream’, ‘Perlita’,‘Planters Jumbo’, ‘PMR 5’, WMR 29 and breeding linePMR Honeydew).

Among a broad germplasm collection of melonsinoculated with MNSV (532 accessions), Pitrat et al.(1996) found 7% immune accessions. The resistance wasconfirmed to be quite common in American cantaloupecultivars (22 resistant accessions representing 28 % outthe North American accessions tested). Some resistantaccessions were found originating also from Far Eastand India. One recessive gene, nsv, controls the resis-tance to MNSV (Coudriet et al., 1981). First described inthe American cultivar Gulfstream, the same gene wasshown to be present in other American germplasm (‘PMR5’, ‘Planters Jumbo’, VA 435) and the Asian accession PI161375 (Coudriet et al., 1981; Pitrat, 1991). nsv wasmapped on the linkage group XII (formely 7) (Pitrat, 1991;Baudracco-Arnas and Pitrat, 1996; Perin et al., 2002).The fine mapping and the cloning of the gene revealedthat the resistance corresponds to a single nucleotide

substitution in the translation initiation factor eIF4E(Morales et al., 2002; Morales et al., 2005; Nieto et al.,2006). The same substitution was found in all the MNSVresistant accessions, suggesting that the resistance hasa unique origin (Nieto et al., 2007).

Two independent dominant genes, named Mnr-1and Mnr-2, were reported to control resistance to sys-temic infection of MNSV in Doublon; Mnr-1 is linked tonsv at 19 cM (Mallor et al., 2003).

No complete sources of resistance to Squash mosaicvirus (SqMV, Comovirus) have been reported in melons.Tolerance was, however, observed in accessions origi-nating from India, Afghanistan, China and Pakistan(Webb and Bohn, 1962; Provvidenti, 1989, 1993). TheKorean and Chinese accessions PI 161375 and China 51(var. makuwa) were described to develop delayed mosaicsymptoms, reduced virus multiplication, and, interest-ingly, complete resistance to seed transmission of SqMV(Maestro-Tejada, 1992; Provvidenti, 1998). Resistanceto seed transmission was shown to be effective againstfour different strains of SqMV (Provvidenti, 1998). Tol-erance to foliar symptoms incited by a melon strain ofSqMV was shown to be controlled by a single recessivegene in China 51, but appeared to be partially dominantagainst a squash pathotype of SqMV (Provvidenti, 1998).Unnamed so far, we propose to name the gene sqmv.

Partial resistance (restriction to the virus move-ment) to the SH isolate of Cucumber green mottle mosaicvirus (CGMMV, Tobamovirus) was reported in themakuwa type Chang Bougi accession (Sugiyama et al.,2006). The resistance was controlled by two complemen-tary, recessive genes, called cgmmv-1 and cgmmv-2(Sugiyama et al., 2007).

Resistance to a complex of viruses from Egypt in PI378062 was reported to be controlled by a single domi-nant gene, named Imy, Interveinal mottling and yellowingresistance (Hassan et al., 1998).

Insect resistanceResistance to the melon-cotton aphid, Aphis gossypii

(Homoptera: Aphididae), was first reported by Kishabaand Bohn. A dominant gene, Ag, was reported to controlantixenosis, antibiosis under controlled no-choice testsand free-curling tolerance in LJ 90634, later called PI414723 (Kishaba et al., 1971, 1976). Pitrat and Lecoq(1980; 1986) reported resistance in PI 161375 and in PI414723 to several viruses when they are transmitted byA. gossypii. The resistance is vector-specific (only A.gossypii), and non-specific to viruses (CMV, ZYMV,WMV…). It co-segregates with antixenosis describedpreviously. Resistance to viruses when they are trans-mitted by A. gossypii, is controlled by a single gene,named Vat (Virus aphid transmission). The Vat locus was


mapped to a subtelomeric position on the linkage groupV (formely 2) (Pitrat, 1991; Baudracco-Arnas and Pitrat,1996; Brotman et al., 2002; Perin et al., 2002). A singlegene was cloned by positional cloning, which confersboth aphid resistance and virus resistance when theyare transmitted by A. gossypii. The gene was shown toencode a CC-NBS-LRR protein (Dogimont et al., 2004;Pauquet et al., 2004; Dogimont et al., 2010). Four addi-tive and two couples of epistatic QTLs affectingbehaviour and biotic potential of A. gossypii weremapped in recombinant inbred lines derived from thecross Védrantais x PI 161375; amongst them, a majorQTL, which affects both behavior and biotic potential ofA. gossypii, corresponds to the Vat gene (Boissot et al.,2010).

A single dominant gene, named Lt, was reportedto control resistance to the leafminer Liriomyza trifolii(Diptera : Agromyzidae) in the old French cultivarNantais Oblong (Dogimont et al., 1999). Resistant plantsexhibit fewer mines and a very high larval mortality.The resistance is inefficient towards L. huidobrensis.

Two complementary recessive genes (dc-1 and dc-2) for resistance to the melon fruit fly, Bractocera cucurbitae(formely Dacus cucurbitae, Diptera: Tephritidae) were re-ported by (Sambandam and Chelliah, 1972).

A monogenic recessive resistance to cucumberbeetles was reported in C922-174-B in crosses amongnon-bitter genotypes. The gene named cbl (=cb) wasshown to be efficient towards three species of Coleoptera:the banded beetle Diabrotica balteata, the spotted beetleD. undecimpunctata howardi and the stripped beetleAcalymna vittatum (Nugent et al., 1984). In AR Top Mark,resistance to D. undecimpunctata howardi was also re-ported to be recessive and linked to the bitterness trait,controlled by the dominant gene Bi (Lee and Janick, 1978)that makes the melon attractive to the spotted beetle(Nugent et al., 1984).

A dominant gene, named Af, was reported to con-trol resistance to the red pumpkin beetle (Aulacophorafoveicollis, Coleoptera: Chrysomelidae) in Casaba(Vashistha and Choudhury, 1974).

Fungal DiseasesFusarium wilt resistance. Three genes were reported

to control resistance to Fusarium oxysporum f.sp. melonis.A single dominant gene, Fom-1, controls resistance to F.oxysporum races 0 and 2; it was reported in the old Frenchcultivar Doublon (Risser, 1973; Risser et al., 1976). Fom-1 was mapped at a distal end of the linkage group IX(formely 5), at 2 cM from the PRSV resistance gene, Prv²(Perin et al., 2002). Molecular markers for Fom-1, usefulfor marker assisted selection, were developed (Brotman

et al., 2005; Oumouloud et al., 2008; Tezuka et al., 2009;Tezuka et al., 2011). A single dominant gene, Fom-2, con-trols resistance to F. oxysporum races 0 and 1; it was re-ported in CM17187 (Risser, 1973; Risser et al., 1976).Fom-2 was mapped to the linkage group XI (Perin et al.,2002). The gene Fom-2 was cloned and reported to en-code a NBS-LRR type R protein of the non-TIR subfam-ily (Joobeur et al., 2004). Molecular markers linked toFom-2 were developed (Zheng et al., 1999; Zheng andWolff, 2000), but their use was not completely satisfyingbecause of recombination (Sensoy et al., 2007). New prom-ising molecular markers were recently designed withinthe gene (Wang et al., 2011). Resistance to F. oxysporumraces 0, 1 and 2 is quite frequent (Alvarez et al., 2005).The Fom-3 gene was reported in Perlita FR; it confers thesame phenotype as Fom-1 but segregates independentlyfrom Fom-1 (Zink and Gubler, 1985).

Resistance to F. oxysporum races 0 and 2 in the Span-ish var. cantalupensis accession Tortuga was reported tobe controlled by two independent genes, one dominantand the other one recessive. The dominant likely is Fom-1; the recessive one was named fom-4 (Oumouloud et al.,2010).

A major recessive gene, named fom1.2a, was re-ported to confer resistance to F. oxysporum race 1.2 in theIsraeli breeding line BIZ. The gene was located at a dis-tal end of the LG II (opposite to the gene a ,andromonoecious) (Herman et al., 2008). A second reces-sive gene was previously reported to segregate in thesame population (Herman and Perl-Treves, 2007). Incontrast, nine QTLs were reported to control the reces-sive resistance to race 1.2 in the French breeding lineIsabelle, derived from the Far East resistant accessionOgon 9 (Perchepied and Pitrat, 2004; Perchepied et al.,2005). The resistance of the var. cantalupensis accessionBG-5384 from Portugal to F. oxysporum race 1.2 (Ypathotype) was also reported to be polygenic and reces-sive (Chikh-Rouhou et al., 2008; Chikh-Rouhou et al.,2010).

Powdery mildew resistance. Several dominant resis-tance genes to powdery mildew were reported in melon.Genetic relationship between these genes is still con-fused, as is the definition of powdery mildew races(McCreight, 2006; Lebeda et al., 2011). Mapping of pow-dery resistance genes and QTLs in several crosses hasthus far located them in six distinct melon linkagegroups.

Jagger et al. (1938) reported a dominant resistancegene, Pm-1, to powdery mildew in ‘PMR 45’. In the origi-nal paper, Pm-1 was reported to confer resistance toErysiphe cichoracearum but the pathogen wasmisidentified and was later determined to have beenPodosphaera xanthii. Pm-1 likely corresponds to the gene


Pm-A, which confers resistance to P. xanthii race 1 in‘PMR 45’, described in Epinat et al. (1993). The powderymildew resistance gene from ‘PMR 45’, introgressed intoa yellow-fleshed breeding line, was reported to be lo-cated in the linkage group IX, loosely linked to the PRSVresistance gene, Prv (Teixeira et al., 2008).

A single dominant gene, Pm-x, confers resistanceto P. xanthii race 1 and 2 (at least) in PI 414723; it waslocated in the linkage group II, linked to the ZYMV re-sistance gene Zym and to the andromonoecious gene a(Pitrat, 1991; Perin et al., 2002).

A single dominant gene was reported in WMR 29,Pm-w, which confers resistance to P. xanthii races 1, 2and 3 (Pitrat, 1991). It likely corresponds to Pm-B inEpinat et al. (1993). It was located in the linkage group V(formerly 2), closely linked to the Vat locus (Pitrat, 1991;Perin et al., 2002).

Harwood and Markarian (1968) reported two domi-nant genes in PI 124112, Pm-4 and Pm-5. These genesmay correspond to the two genes of PI 124112 reportedin Perchepied et al. (2005), PmV.1 and PmXII.1. PmV.1confers resistance to P. xanthii races 1, 2, and 3 and waslocated in the linkage group V, closely linked to the Vatlocus. Pm-XII.1 confers resistance to P. xanthii races 1, 2and 5 and to Golovinomyces cichoracearum race 1 and wasmapped to the linkage group XII. It may correspond toone of the two genes, Pm-F and Pm-G, which were re-ported to interact for controlling resistance to G.cichoracearum in PI 124112 (Epinat et al., 1993).

Two genes were reported in ‘PMR 5’, Pm-1 andPm-2 (Bohn and Whitaker, 1964). Allelism tests clearlyshowed that ‘PMR 5’ has the same gene as ‘PMR 45’ tocontrol P. xanthii race 1. Pm-2 likely corresponds to Pm-C, which confers resistance to P. xanthii race 2 in inter-action with Pm-1 (Epinat et al., 1993). Two genes, Pm-C(Pm-2) and Pm-E, were suggested to interact in ‘PMR 5’to control resistance to G. cichoracearum (Epinat et al.,1993). Recently, two QTLs of resistance to P. xanthii race1 and N1 were located in the linkage groups II and XII inrecombinant inbred lines derived from the cross PMARNo.5 x Harukei No.3 (Fukino et al., 2006; Fukino et al.,2008). These two QTLs may correspond to the same ge-nomic regions as reported in PI 124112, with differentalleles. PMAR No.5 (= AR 5) was obtained from an aphidresistant line and successive backcrosses to ‘PMR 5’(McCreight et al., 1984). The results obtained by (Fukinoet al., 2006; Fukino et al., 2008) suggest that powderymildew resistance genes in PMAR No.5 may be differentfrom those in ‘PMR 5’, as Pm-1 is expected to be locatedin the linkage group IX (Teixeira et al., 2008).

Harwood and Markarian (1968) reported a singledominant resistance gene in PI 124111, Pm-3. Kenigsbuch

and Cohen (1989) reported a second gene in PI 124111,Pm-6, independent from Pm-3, which confers resistanceto P. xanthii race 2. Their relationship with the otherpowdery mildew resistance genes is unknown.

Resistance to the Chinese race of P. xanthi (with aunique reaction pattern of the commonly used melonrace differentials) in the Indian accession PI 134198 wasreported to be controlled by a single dominant gene, des-ignated Pm-8, which was suggested to be located in thelinkage group VII (Liu et al., 2010).

Resistance to P. xanthi races 1, 2 and 5 in TGR-1551 was reported to be controlled by two independentgenes, one dominant and one recessive, each one con-ferring resistance to all three races (Gómez-Guillamónet al., 2006; Yuste-Lisbona et al., 2008). The dominantgene, Pm-R, was recently located in the linkage group V,closely linked to the Vat and Pm-w loci (Yuste-Lisbona etal., 2011); the recessive gene was putatively located inthe linkage group VIII, with a LOD score lower than thethreshold score (Yuste-Lisbona et al., 2011).

In the same manner, resistance to P. xanthii in PI313970 or 90625 was reported to be controlled by domi-nant, co-dominant, and recessive genes (McCreight,2003; McCreight and Coffey, 2007; Pitrat and Besombes,2008). Recently, PI 313970 resistance to the race S, a newstrain of P. xanthii from Eastern-USA, virulent on all thecommonly used resistance differentials, was reported tobe controlled by a single recessive gene, named pm-S.The relationship of pm-S with the previously reportedresistance genes in PI 313970 is unknown (McCreightand Coffey, 2011).

Other fungi. Several genes have been described tocontrol resistance to gummy stem blight, caused byDidymella bryoniae (asexual form Mycosphaerellacitrullina). Four independent dominant genes, Gsb-1through Gsb-4, were reported to confer a high level ofresistance in PI 140471, PI 157082, PI 511890, and PI482398 (Prasad and Norton, 1967; Frantz and Jahn,2004). In the latter accession, a recessive gene, gsb-5, in-dependent from Gsb-1, Gsb-2, Gsb-3 and Gsb-4 was alsoreported (Frantz and Jahn, 2004). A single dominant gene(previously named Mc-2), was reported to confer a mod-erate level of resistance in C-1 and C-8 (Prasad andNorton, 1967); we propose to rename it Gsb-6.

A single dominant gene, Ac, was reported to con-trol resistance to Alternaria cucumerina in the line MR-1(Thomas et al., 1990). A semi-dominant gene, Mvd, wasreported to control partial resistance to melon vine de-cline caused by Acremonium cucurbitacearum andMonosporascus cannonballus in the wild type accessionPat 81 (Iglesias et al., 2000).


Oomycete resistanceSources of resistance to downy mildew caused by

the oomycete Pseudoperonospora cubensis were reportedin several Indian accessions (Dhillon et al., 2007; Ferganyet al., 2011). Downy mildew resistance was reported tobe controlled by two partially dominant, complemen-tary genes, Pc-1 and Pc-2, in the Indian accession PI124111 (Cohen et al., 1985; Thomas et al., 1988;Kenigsbuch and Cohen, 1992). This accession was re-ported to be resistant to the six known pathotypes ofdowny mildew (Cohen et al., 2003). Two complemen-tary, dominant genes (Pc-4 and Pc-1 or Pc-2) were alsoreported to control resistance to downy mildew in an-other Indian accession PI 124112 (Kenigsbuch andCohen, 1992). Nine QTLs for resistance to P. cubensiswere located on a melon map developed from the cross‘Védrantais’ x PI 124112. Among them, a major QTL, Pc-XII.1, was located in the linkage group XII, closely linkedto the powdery mildew resistance QTL Pm-XII.1, whichconfers resistance to P. xanthii races 1, 2 and 5 and G.cichoracearum race 1 (Perchepied et al., 2005). A singledominant gene of partial resistance, Pc-3, was reportedin the Indian accession PI 414723 (Epinat and Pitrat,1989). The gene Pc-5 was reported to interact with themodifier gene, M-Pc-, to control downy mildew resis-tance in the line 5-4-2-1; in presence of M-Pc-5, the resis-tance conferred by the gene Pc-5 is dominant, while inabsence of M-Pc-5, the resistance is recessive (Angelovand Krasteva, 2000).

Seed and Seedling GenesThree genes were reported to control seed coat

color: the r gene (red stem) controls brown seed color anda red stem in PI 157083 (30569) (Bohn, 1968; McCreightand Bohn, 1979). The gene Wt (White testa) controls whiteseed testa color and is dominant to yellow or tan seedcoat color (Hagiwara and Kamimura, 1936). A White testagene (Wt-2) was also reported in PI 414723, dominant toyellow seed testa color and mapped to the linkage groupIV (Périn et al., 1999). The pine-seed shape of the seedsof PI 161375 is controlled by a single recessive gene, pin,pine-seed shape, which was mapped to the linkage groupIII (Perin et al., 2002). This trait is common in melon inthe pinonet Spanish type. The presence of a gelatinoussheath around the seeds (versus absence) was reportedto be controlled by a single dominant gene Gs, Gelatinoussheath (Ganesan, 1988).

Several chlorophyll deficient mutants were re-ported in melon. A single recessive gene, alb, (albino) con-trols the white cotyledon, lethal mutant in Trystorp(Besombes et al., 1999). The dominant pale cotyledonsmutant Pa, Pale, is a lethal mutation as PaPa are albinos

and die early, while PaPa+ have yellow cotyledons andleaves (McCreight and Bohn, 1979); Pa was shown to belinked to the gl (glabrous) and r (red stem) mutant genes(Pitrat, 1991). A single recessive gene, yg (yellow green),controls light green cotyledons and leaves in the line26231 (Whitaker, 1952); it was located in the linkagegroup XI (former 6) (Pitrat, 1991). An allele of yg, firstdescribed as lg (light green) in the cross Dulce x TAM-Uvalde, was renamed ygw (yellow green Weslaco) (Cox,1985; Cox and Harding, 1986). A single recessive gene, f(flava), controls bronze yellow cotyledons and leavesand a reduced plant growth in the Chinese accessionK2005 (Pitrat et al., 1986); it was reported to be closelylinked to the lmi (long main stem internode) gene (Pitrat,1991). A recessive mutant with a yellow ring on the coty-ledons that later disappears, leaving the plants a nor-mal green, was named h (halo) (Nugent and Hoffman,1974); it was shown to be linked to the genes a(andromonoecious), Pm-x (Powdery mildew resistance x) andZym (Zucchini yellow mosaic virus resistance) and was thenlocated in the linkage group II (former 4) (Pitrat, 1991;Perin et al., 2002). Three recessive virescent genes v, v-2and v-3 control pale cream cotyledons and hypocotyls,which turn green later; the younger leaves are light greenwhile the older ones are normal green (Hoffman andNugent, 1973; Dyutin, 1979; Pitrat et al., 1995); the v-3gene was shown to be independent to v (Pitrat et al.,1995). Two yellow virescent recessive mutant genes, yv(yellow virescent) and yv-2, were reported (allelism un-known); they control pale cotyledons, yellow green youngleaves and tendrils and green older leaves, associatedwith a severely reduced plant growth (Zink, 1977; Pitratet al., 1991).

The incapacity of a mutant to efficiently absorb Fe(iron) and Mn (manganese) was reported to be controlledby a recessive gene, fe; the mutant chlorotic leaves withgreen veins turn to green when iron is added to the nutri-ent solution (Nugent and Bhella, 1988; Jolley et al., 1991).

A single recessive gene, ech (exaggerated curvatureof the hook), was shown to control the triple response ofseedling germination in the dark in the presence of eth-ylene. Seedlings exhibit a very strong, 360° hook curva-ture of hypocotyls in PI 161375 (ech), while they exhibit amoderate, 180° curvature in ‘Védrantais’ and PI 414723(Ech). The ech gene was mapped to the linkage group I(Perin et al., 2002).

Seedling bitterness due to the presence ofcucurbitacins, common in honeydew or Charentais type,was shown to be dominant over non-bitter, found in mostAmerican cantaloupes, and controlled by a single geneBi (Bitter) (Lee and Janick, 1978).

A single recessive delayed lethal mutant, dlet (for-merly dl) was described by Zink (1990); it exhibits a re-


duced growth, necrotic lesions on leaves leading to pre-mature death.

Leaf and Foliage GenesSeveral genes control leaf and foliage traits in

melon. Two linked dominant genes, Ala (Acute leaf apex)and L (Lobed leaves) were reported to control leaf shapein ‘Main Rock’ (Ala and L) crossed with ‘PV Green’ (alaand l) (Ganesan and Sambandam, 1985). Highly in-dented leaves, instead of round, are controlled by a singlerecessive gene, dl (dissected leaf), in URRS 4 (Dyutin,1967). An allele of dl in ‘Cantaloup de Bellegarde’, pre-viously described as cut leaf, was named dlv, dissected leafVelich (Velich and Fulop, 1970). A second gene dl-2 (dis-sected leaf-2), allelism unknown, was reported as “hojashendidas” (Esquinas Alcazar, 1975). A single dominantgene, Sfl, was reported to control the subtended floral leaftrait; the leaves bearing hermaphrodite/pistillate flow-ers in their axis, are sessile, small and enclosing the flow-ers in ‘Makuwa’, Sfl, while normal in ‘Annamalai’, sfl(Ganesan and Sambandam, 1979). Cox (1985) reportedtwo recessive leaf mutant genes, brittle leaf dwarf (bd) andcurled leaf (cl), which both affect the female fertility. Spoon-shaped leaves with upward curling of the leaf marginswere reported to be controlled by a single recessive gene,named cf (cochleare folium) in a spontaneous mutant in‘Galia’ (Lecouviour et al., 1995). A single recessive gene,gl (glabrous), was reported to control completely hairlessplants in Arizona glA (Foster, 1963). A single recessivegene, r (red stem), controls in PI 157083 (30569) a redstriped hypocotyl and red stem, especially at internodes,that is photosensitive, and reddish or tan seed coat color(Bohn, 1968; McCreight and Bohn, 1979). The genes gland r were shown to be linked in a same linkage group(LG 3) comprising also Pa (Pale) and ms-1 (male sterile-1)(McCreight, 1983; Pitrat, 1991).

Plant Architecture GenesA single gene, recessive or incompletely dominant,

called slb, short lateral branching (formerly sb) was sug-gested to control the short lateral branching trait in LB-1, a wild melon from Russia (Ohara et al., 2001). In 2008,(Fukino et al., 2008) reported two QTL for short lateralbranching in a cross between a breeding line Nou 4 de-rived from LB-1 and the normal branching ‘Earl’sFavourite’ (Harukei 3). The QTL mapped to LG VII andLG XI, explained, respectively, 14.8 % (The allele ofHarukei 3 contributed to shorter length branches) and42.2% (The allele of ‘Nou 4’ contributed to shorter lengthbranches). A mutant lacking lateral branches, named ab,abrachiate, was reported; it produces only male flowers(Foster and Bond, 1967).

A single recessive gene, lmi (long main-stem intern-ode), controls a long hypocotyl and a long internodelength (about 20 cm) in the main stem but does not affectinternode length of lateral branches in 48764(McCreight, 1983). Three recessive genes that controlledshort-internodes, si-1, si-2, si-3 (short internode-1, -2, -3),were reported in three independent melon lines, UCTopmark Bush, Persia 202, and ‘Maindwarf’ (Denna,1962; Paris et al., 1984; Knavel, 1990). si-1 plants dis-play a bush phenotype, with an extremely compact grow-ing habit and very short (about 1 cm) internode length(Denna, 1962; Zink, 1977); si-1 is linked to the gene yv,yellow virescent (Pitrat, 1991). Internodes of si-2 and si-3plants are short but less compact than si-1 plants. In si-2 plants, the first internodes are short, leading to a ‘bird’snest’ phenotype; later internodes are not modified. In si-3 plants, internode length is reduced at all plant devel-opment stages. Fasciation of the main stem (reaching upto 15 cm) in the Charentais type line Vilmorin 104 wascontrolled by a single recessive gene, named fas, fasci-nated (Gabillard and Pitrat, 1988).

Flower GenesSex determination in melon is controlled by two

major genes, a and g. The andromonoecious gene a (Rosa,1928; Poole and Grimball, 1939; Wall, 1967) controls themonoecious versus andromonoecious sex type in melon.The gene mapped to the linkage group II (Perin et al.,2002; Silberstein et al., 2003). The gene was recentlycloned and was shown to encode an ACC synthase gene,CmACS7. The transition between monoecy andandromonoecy is conferred by a single substitution,which leads to an inactive form of this key enzyme inthe ethylene biosynthesis (Boualem et al., 2008). Molecu-lar markers linked to the gene (Noguera et al., 2005;Sinclair et al., 2006; Kim et al., 2010) and within the geneare available (Boualem et al., 2008).

The gynoecious, g, gene controls the transition ofmonecious plants to gynoecious plants carrying onlyfemale flowers. The gene was mapped to a distal end ofthe linkage group V, opposite to the Vat gene. Positionalcloning of the gene showed that the gene G encodes for atranscription factor of the WIP family, CmWIP1. Thegynoecious allele g corresponds to the insertion of a trans-posable element, which epigenetically represses the ex-pression of CmWIP1 (Martin et al., 2009). A third gene,named gy (gynomonoecious, previously also called n orM), interacts with a and g to produce stable gynoeciousplants in the gynoecious line WI 998 (Kenigsbuch andCohen, 1987, 1990).

Five single recessive genes of male-sterility includ-ing ms-1 to ms-5 were reported in melon (Bohn andWhitaker, 1949; Bohn and Principe, 1964; Lozanov,


1983; McCreight and Elmstrom, 1984; Lecouviour et al.,1990) in (Pitrat, 1991, 2002). Each of these genes dis-plays a unique phenotype. The five sterility genes werelocated in five different linkage groups (Pitrat, 1991; Parket al., 2009). McCreight (1983) and Pitrat (1991) reportedloose linkages between red stem (r) and the ms-1 gene,and between yellow green leaves (yg) and the ms-2 gene,respectively. Park et al. (2009) mapped the ms-3 gene tothe linkage group 9 of the linkage map Deltex x TGR-1551, which corresponds to the linkage group VII.

A Macrocalyx dominant gene, Mca, was reported tocontrol the presence of large, leafy sepals in staminateand hermaphrodite flowers in the Japanese cultivarMakuwa (Ganesan and Sambandam, 1979). Two reces-sive genes were reported to modify the color of petals; gp(green petals) and gyc (greenish yellow corolla) control thepresence of a green corolla with venation or the pres-ence of a greenish yellow corolla, instead of the normalyellow corolla (Mockaitis and Kivilaan, 1965; Zink,1986).

Rosa (1928) reported that tricarpellary ovary wasmonogenically inherited over pentacarpellary ovaryfound in Cassaba melons; the gene, named p (pentamer-ous) was mapped to the linkage group XII, closely linkedto the major QTL for CMV resistance (Dogimont et al.,2000; Perin et al., 2002; Essafi et al., 2009). A single re-cessive gene, n (nectarless), was reported to control theabsence of nectar in all flowers in the mutant 40099 (Bohn,1961).

Fruit GenesFruit shape was reported to be controlled by a single

gene O (Oval shape), dominant to round, and associatedwith andromonoecious gene a (Wall, 1967). As early as1928, Rosa (1928) noted the association of elongate fruitwith pistillate flowers (monoecious plants) and globu-lar fruit with perfect flowers (andromonoecious plants)in segregating populations. More recently, several fruitshape QTL were mapped in several populations to atleast five linkage groups; one of them co-localized withthe a locus on the linkage group II (Perin et al., 2002;Monforte et al., 2004; Eduardo et al., 2007; Fernandez-Silva et al., 2010; Díaz et al., 2011). Spherical fruit shapewas also reported to be controlled by a single gene, sp(spherical fruit shape), recessive to an obtuse fruit shape(Lumsden, 1914; Bains and Kang, 1963); this gene maybe the same as the gene O.

A single dominant gene, Ec (Empty cavity), was re-ported to control the presence of separated carpels atfruit maturity, leaving a cavity in PI 414723 fruit (ec in‘Védrantais’) (Périn et al., 1999). The Ec gene wasmapped to the linkage group III (Perin et al., 2002).

External fruit appearance. Rind color of melon fruitvarieties includes white, yellow, orange, or green, andcan be variegated. The white color of immature fruitswas reported to be dominant to green immature fruitsand controlled by a single gene, Wi, White color of imma-ture fruit (Kubicki, 1962). The white color of mature fruitswas, in contrast, reported to be controlled gene w, white,recessive to dark green fruit skin in a cross betweenHoneydew (w) and Smiths’ Perfect cantaloupe (W, darkgreen) (Hughes, 1948). Melon rind color was shown tobe based on different combinations of three major pig-ments, chlorophyll, carotenoids and naringerin-chal-cone, a flavonoid pigment responsible for the yellow colorof mature fruits in Yellow Canari melon type (Tadmor etal., 2010). Accumulation of naringerin-chalcone was re-ported to be inherited as a monogenic dominant trait inthe cross ‘Noy Amid’ (yellow rind) x ‘Tendral VerdeTardio’ (dark green rind); accumulation of chlorophylland carotenoids segregates jointly as a single dominantgene, independent to naringerin-chalcone accumulation(Tadmor et al., 2010). We propose to name Nca the gene,which regulates naringerin-chalcone accumulation (versusnon-accumulation), and Chl and Car the two linkedgenes, which control chlorophyll and carotenoid accumu-lation in the rind of mature fruit, respectively. In addi-tion, minor genes likely control quantitative variation ofthe accumulation of these pigments. A polygenic con-trol of the external fruit color was reported in the cross‘Piel de Sapo’ x PI 161375 (Whitaker and Davis, 1962;Monforte et al., 2004; Eduardo et al., 2007; Obando et al.,2008).

Vein tracts, formerly and incorrectly referred to assutures, on the fruit rind was reported to be controlledby a single recessive gene s, sutures (Bains and Kang,1963; Davis 1970). The same inheritance was found intwo crosses: ‘Védrantais’ (s-2, presence of sutures) x PI161375 (S-2, without sutures) and ‘Védrantais’ x PI414723 (S-2). The s-2 gene was mapped to the linkagegroup XI (Perin et al., 2002). Stripes on the rind was re-ported to have a monogenenic recessive inheritance(gene st for striped epicarp) by (Hagiwara and Kamimura,1936). The presence of stripes on young fruits of ‘Dulce’(before netting development) was also reported to becontrolled by a single recessive gene, st-2 (striped epi-carp-2), in the cross Dulce (st-2) x PI 414723 (St-2, non-striped) (Danin-Poleg et al., 2002); the gene st-2 wasmapped to the linkage group XI. Further studies wouldbe required to clarify the relationship between st-2 and s-2, also located in the linkage group XI.

The ridge fruit surface was reported to be con-trolled by a single gene, ri (ridge in C68), recessive toridgeless (Ri in ‘Pearl’) (Takada et al., 1975). The speck-led epidermis of the fruit is controlled by a single reces-


sive gene, spk (speckled fruit epidemis) in PI 414723 (Spk in‘Védrantais’) and was mapped to the linkage group VII(Perin et al., 2002). A single gene, Mt (Mottled rind pat-tern), was reported to control a mottled rind in‘Annamalai’, dominant to uniform color mt in ‘Makuwa’(Ganesan, 1988). The presence of dark spots (about 1 cmin diam.) on the rind (versus no spots) has a monogenicrecessive inheritance in crosses Védrantais (Mt-2) x PI161375 (mt-2) and Védrantais (Mt-2) x PI 414723 (mt-2),as the F1 fruits have a uniform color rind (Périn et al.,1999); it was erroneously named Mt-2 in the previousgene list. mt-2 was mapped to the linkage group II (Perinet al., 2002).

A single dominant gene governing the develop-ment of net tissue, regardless of the degree of nettingwas reported in BIZ in a cross with smooth-skinned PI414723 (Herman et al., 2008). We propose to name thegene Rn (Rind netting) instead of N. The gene was mappedto the linkage group II, closely linked to fom1.2a forFusarium wilt resistance; additional minor loci likelyaffect the density of the net (Herman et al., 2008). SeveralQTL for the height and the width of the net in ‘Deltex’were detected in a cross between netted ‘Deltex’ and net-free TGR-1551 (Park et al., 2009).

Melon fruit flesh color has been proposed to becontrolled by two genes, gf for green flesh in Honeydew,recessive to orange flesh (Gf in Smiths’ Perfect canta-loupe) (Hughes, 1948) and wf for white flesh (Iman et al.,1972). Genetic control of melon mesocarp color has, how-ever, not been clearly elucidated and likely differs amongmarket types. Clayberg (1992) confirmed that green andwhite mesocarps are recessive to orange and indicatedthat gf and wf interact epistatically. Mesocarp color (or-ange vs. green) segregated as a single recessive gene inrecombinant inbred lines derived from orange fleshVédrantais x green flesh PI 161375 (Perin et al., 2002)and orange flesh AR 5 x green flesh Harukai N°3(Fukino et al., 2008). The segregating gene, named gf,proposed to be renamed wf, mapped to the linkage groupIX. In F2 and doubled haploid lines derived from thecross between green mesocarp PI 161375 and white me-socarp Piel de Sapo T111, individuals with orange me-socarp were observed at a low frequency (Monforte etal., 2004); a single recessive gene segregated, if orangemesocarp phenotype was excluded and mapped to thelinkage group VIII (formerly G1) (Monforte et al., 2004).Several QTL for fruit flesh color were described in nearisogenic lines derived from the same cross (Eduardo etal., 2007; Obando et al., 2008). Recently, three QTL asso-ciated with color variation (white, green, orange) withputative epistatic interaction were identified in the crossbetween the white-fleshed Chinese line Q3-2-2 and or-ange-fleshed ‘Top Mark’ (Cuevas et al., 2009; Cuevas et

al., 2010). Five QTL associated with beta-carotene con-tent, which is related to color intensity of the mesocarp,were identified in the cross between two orange-fleshedgenotypes, USDA 846-1 and ‘Top Mark’ (Cuevas et al.,2008).

Sweet melon cultivars are characterized by highsucrose and low acid levels in mature fruit flesh. A single,incompletely recessive gene, suc, controlled accumula-tion of sucrose in the cross between the low sucroseFaqqous (var. flexuosus) and the high sucrose ‘NoyYizre’el’ (Burger et al., 2002). Several QTL associatedwith total soluble solid content and sugar content havebeen described in several populations (Monforte et al.,2004; Sinclair et al., 2006; Park et al., 2009; Harel-Beja,2010).

A dominant gene, So (Sour) was reported to controlhigh acidity in melon fruit (Kubicki, 1962). A single domi-nant gene, So-2 (Sour-2) for sour taste of the mature fruit,was also reported in PI 414723 (Périn et al., 1999; Burgeret al., 2003). A single recessive gene, pH, was reported tocontrol fruit flesh acidity in PI 414723. Low pH value inPI 4141723 was dominant to high pH value in ‘Dulce’.The pH gene was mapped to the linkage group VIII(Danin-Poleg et al., 2002); it likely corresponds to So-2.

While ripe melon fruits usually do not have a bit-ter taste, young fruits are divided into two types: bitterand non-bitter. A single dominant gene, Bif-1 (Bitter fruit-1, formely Bif), was reported to control the strong bittertaste of tender fruits in Indian wild melon (Parthasarathyand Sambandam, 1981). A monogenic dominant inher-itance for the bitterness of young fruits was confirmedin wild melons from Africa and China (Ma et al., 1997).The cross of non-bitter melon lines (var. conomon andvar.makuwa) with var. inodorus and var. cantalupensis)yielded, however, bitter young melons, which suggestscomplementary gene action of two independent genes,Bif-2 and Bif-3 (Bif-2_ Bif-3_ are bitter; bif-2bif-2 Bif-3_and Bif-2_ bif-3bif-3 are non-bitter) (Ma et al., 1997). Oneof them may be the same as Bif-1. The relationship withthe gene Bi controlling seedling bitterness (Lee andJanick, 1978) is unknown.

While the single dominant gene Mealy, Me, wasreported to control mealy flesh texture by Ganesan (1988)in an accession named C. callosus crossed with a crisp-fleshed ‘Makuwa’, a monogenic recessive inheritancewas found for the mealy flesh texture in the var.momordica accession PI 414723 (me-2) crossed by‘Védrantais’ (Me-2) (Périn et al., 1999) (included errone-ously as Me-2 in the previous gene list, it is now in-cluded as me-2). A monogenic recessive inheritance wasreported for the juicy character of melon fruit flesh; thegene was named juicy flesh, symbolized jf (Chadha etal., 1972). A single gene was reported to control the


musky flavor of C. melo callosus (Mu, Musky), dominantto the mild flavor in ‘Makuwa’ or ‘Annamalai’ (mu)(Ganesan, 1988).

Fruit abscission at maturity was reported to be con-trolled by two independent loci in two independent stud-ies. In absence of allelism tests, the genes were namedabscission layer Al-1 and Al-2 in C68, al-1 and al-2 in‘Pearl’ (Takada et al., 1975), and Al-3 and Al-4 in theclimacteric Charentais type ‘Védrantais’ (Perin et al.,2002). Al-3 and Al-4 were mapped to the linkage groupsVIII and IX in a recombinant inbred population derivedfrom a cross between ‘Védrantais’ and the non-climac-teric PI 161375 (Perin et al., 2002). A single dominant

gene, Al-5, was reported to control fruit abscission layerformation in the climacteric western shipper type ‘TAMUvalde’ in the cross with the non-climacteric Casabatype ‘TAM Yellow Canary’ (Zheng et al., 2002).

Organogenic competence varies among melongenotypes. In vitro shoot regeneration capacity was re-ported to be controlled by two independent genes, par-tially dominant, Org-1 and Org-2 (Organogenic response)(Molina and Nuez, 1996). A single dominant gene, Org-3, was reported to control the high regeneration compe-tence in the line BU-21/3, in crosses with the low regen-eration competent lines ‘PMR 45’ and ‘AnanasYokneam’ (Galperin et al., 2003).


Table 1. Reported host plant resistance and morphological genes of melon, including genessymbol, synonyms, descriptions, and linkage groups.z


Table 2. Reported melon isozyme genes, including gene symbol, synonym, description,location on the melon genome and reference.

Gene symbol Prefered Synonym Gene description and type lines LG

z References

Aco-1 Ac Aconitase-1. Isozyme variant with two alleles, each regulating one band, in PI 218071, PI 224769.

A (Staub et al., 1998)

Acp-1 APS-11, Ap-11

Acid phosphatase-1. Isozyme variant with two codominant alleles, each regulating one band. The heterozygote has two bands.

(Esquinas Alcazar, 1981)

Acp-2 Acp-1 Acid phosphatase-2. Isozyme variant with two alleles, each regulating one band, in PI 194057, PI 224786. Relationship with Acp-1 is unknown.

(Staub et al., 1998)

Acp-4 - Acid phosphatase-4. Isozyme variant with two alleles, each regulating one band, in PI 183256, PI 224786. Relationship with Acp-1 unknown, different from Acp-2.


Ak-4 - Adenylate kinase. Isozyme variant with two alleles, each regulating one band, in PI 169334.


Fdp-1 - Fructose diphosphate-1. Isozyme variant with two alleles, each regulating one band, in PI 218071, PI 224688.


Fdp-2 - Fructose diphosphate-2. Isozyme variant with two alleles, each regulating one band, in PI 204691, PI 183256.


Gpi - Glucosephosphate isomerase. Isozyme variant with two alleles, each regulating one band, in PI 179680.


Idh - Isocitrate dehydrogenase. Isozyme variant with two alleles, each regulating one band, in PI 218070, PI 224688.


Mdh-2 - Malate dehydrogenase-2. Isozyme variant with two alleles, each regulating one band, in PI 224688, PI 224769.

B (Staub et al., 1998)







Mpi-1 - Mannosephosphate isomerase-1. Isozyme variant with two alleles, each regulating one band, in PI 183257, PI 204691.


Mpi-2 - Mannosephosphate isomerase-2. Isozyme variant with two alleles, each regulating one band, in PI 183257, PI 204691.


Pep-gl - Peptidase with glycyl-leucine. Isozyme variant with two alleles, each regulating one band, in PI 218070.


Pep-la - Peptidase with leucyl-alanine. Isozyme variant with two alleles, each regulating one band, in PI 183256.


Pep-pap - Peptidase with phenylalanyl-proline. Isozyme variant with two alleles, each regulating one band, in PI 183256.


Pgd-1 6-PGDH-21 Pgd-21

Phosphoglucodehydrogenase-1. Isozyme variant with two alleles, each regulating one band.The heterozygote has one intermediate band.


6-Pgd-2 - 6-Phosphogluconate dehydrogenase. Isozyme variant with two alleles, each regulating one band, in PI 161375, Védrantais. Relationship with Pgd-1 is unknown.

IX (Baudracco-Arnas and Pitrat, 1996)


Gene symbol Prefered Synonym Gene description and type lines LG

z References

Pgd-3 Pgd 6-Phosphogluconate dehydrogenase. Isozyme variant with two alleles, each regulating one band, in PI 218070. Relationship with Pgd-1 and 6-Pgd-2 is unknown.


Pgi-1 PGI-11 Phosphoglucoisomerase-1. Isozyme variant with two alleles, each regulating two bands. The heterozygote has three bands.


Pgi-2 PGI-21 Phosphoglucoisomerase-2. Isozyme variant with two alleles, each regulating two bands. The heterozygote has three bands.


Pgm-1 PGM-21 Pgm-21

Phosphoglucomutase-1. Isozyme variant with two alleles, each regulating two bands. The heterozygotes has three bands.


Pgm-2 Pgm Phosphoglucomutase. Isozyme variant with two alleles, each regulating one band, in PI 218070, PI 179923. Relationship with Pgm-1 is unknown.


Px-1 PRX-11 Peroxidase-1. Isozyme variant with two codominant alleles, each regulating a cluster of four adjacent bands. The heterozygote has five bands.


Px-2 Px2A Prx2

Peroxidase-2. Isozyme variant with two codominant alleles, each regulating a cluster of three adjacent bands. The heterozygote has four bands.

(Dane, 1983; Chen et al., 1990)

Skdh-1 - Shikimate dehydrogenase-1. Isozyme variant with two codominant alleles, each regulating one band. The heterozygote has three bands.

Chen et al., 1990) (Gang and Lee, 1998)

zLinkage groups to which the genes belong are indicated as letters, according to Staub et al. (1998), and Roman numbers

according to Périn et al. (2002).


Table 3. Quantitative traits loci, including description of the quantitative trait, number ofQTL reported, parental lines of the cross used, and references.

,

Description of the quantitative trait, parental lines of the cross used

References

Aphis gossypii resistance Four additive and two couples of epistatic QTL affecting behaviour and biotic potential of Aphis gossypii in the cross Védrantais x PI 161375 (RILs).

(Boissot et al., 2010)

Bemisia tabaci resistance Two QTL affecting the biotic potential of the whiteflies in the cross Védrantais x PI 161375 (RILs).

(Boissot et al., 2010)

Cucumber mosaic virus resistance. Seven QTL are involved in resistance to three different CMV strains in the cross Védrantais x PI 161375 (RILs). A single QTL required for controlling CMV P9 and P104.82 strains in the cross Piel de Sapo x PI 161375 (LG XII).

(Dogimont et al., 2000) (Essafi et al., 2009)

Fusarium oxysporum f.sp. melonis race 1.2 resistance Nine QTL described in the cross Védrantais x Isabelle.

(Perchepied et al., 2005)

Pseudoperonospora cubensis resistance Nine QTL for resistance to downy mildew described in the cross Védrantais x PI 124112.

(Perchepied et al., 2005)

Podosphaera xanthii resistance Two QTL for resistance to powdery mildew described in the cross TGR-1551 x Bola de Ora (F2), a major one, dominant (LG V) and a minor one, recessive (LG VIII).

(Yuste-Lisbona et al., 2011)

Ovary shape Six QTL for ovary length, eight QTL for ovary width and six QTL for the ratio ovary length/ovary width described in the cross Védrantais x PI 161375 (RILs). Five QTL for ovary shape in the cross Piel de Sapo x PI 161375 (NILs).

(Perin et al., 2002) (Eduardo et al., 2007)

Fruit shape Four QTL for fruit length, 5 QTL for fruit width and 6 QTL for the ratio fruit length/fruit width described in the cross Védrantais x PI 161375. Four QTL for fruit length, one for fruit width and two for the ratio fruit length : fruit width described in the cross Védrantais x PI 414723, which are common to both crosses. Eight QTL for fruit shape described in the cross Piel de Sapo x PI 161375 (F2 and DHLs). Eleven QTL for fruit length, 10 QTL for fruit width and 15 QTL for the ratio fruit length/fruit width described in the cross Piel de Sapo x PI 161375 (NILs). Two QTL for fruit length, 2 QTL for fruit width and QTL for the ratio fruit length/fruit width described in the PI 414723 x Dulce (RI).

(Perin et al., 2002) (Perin et al., 2002) (Monforte et al., 2004) (Eduardo et al., 2007; (Fernandez-Silva et al., 2010) (Harel-Beja et al., 2010)

Fruit weight Six QTL described in the cross Piel de Sapo x PI 161375(F2 and DHLs). Eleven QTL described in the cross Piel de Sapo x PI 161375 (NILs).

(Monforte et al., 2004) (Eduardo et al., 2007)

Fruit firmness Two QTL for fruit firmness of the whole fruit described in the cross PI 414723 x Dulce (RI).

(Harel-Beja et al., 2010)

Rind traits Three QTL for stripes, three QTL for sutures described in the cross PI 414723 x Dulce (RI).

(Harel-Beja et al., 2010)

External color of the fruit Four QTL described in the cross Piel de Sapo x PI 161375 (F2 and DHLs). Four QTL described in the cross Piel de Sapo x PI 161375 (NILs). Thirteen QTL for skin color and 12 QTL for ground spot color using the three color components in the cross Piel de Sapo x PI 161375 (NILs).

(Monforte et al., 2004) (Eduardo et al., 2007) (Obando et al., 2008)

Flesh color Three QTL for orange flesh color described in the cross Piel de Sapo x PI 161375 (F2 and DHLs). Four QTL for fruit flesh color described in the cross Piel de Sapo x PI 161375 (NILs). Sixteen QTL for flesh color and 10 QTL for juice color using the three color components in the cross Piel de Sapo x PI 161375 (NILs). Three QTL for flesh color described in the cross PI 414723 x Dulce (RI).

(Monforte et al., 2004) (Eduardo et al., 2007) (Obando et al., 2008) (Harel-Beja et al., 2010)


Description of the quantitative trait, parental lines of the cross used

References

Sugar content of fruit flesh in mature fruit Five QTL for soluble solid content described in the cross Piel de Sapo x PI 161375 (F2 and DHLs). QTL for sucrose, total soluble solids in the cross TAM Dulce x TGR-1551 (F2). Fifteen QTL for soluble solid content in the cross Piel de Sapo x PI 161375 (NILs). Twenty-seven QTL for sugars, eight for fructose, six for glucose, four for sucrose, nine for sucrose equivalents in the cross Piel de Sapo x PI 161375 (NILs). Six QTL for sucrose, total soluble solids in the cross Deltex x TGR-1551 (F2). Six QTL for sucrose, total soluble solids in the cross PI 414723 x Dulce (RI).

(Monforte et al., 2004) (Sinclair et al., 2006) (Eduardo et al., 2007) (Obando-Ulloa et al., 2009) (Park et al., 2009) (Harel-Beja et al., 2010)

Organic acid profile of fruit flesh in mature fruit Twenty-one QTL for organic acids in the cross Piel de Sapo x PI 161375 (NILs).

(Obando-Ulloa et al., 2009)

Ascorbic acid One QTL in the in the cross Deltex x TGR-1551 (F2).

(Park et al., 2009)

Ethylene production in fruit (climacteric crisis). Four QTL described in the cross Védrantais x PI 161375 (RILs). One QTL for ethylene production and climacteric response in the cross Piel de Sapo x PI 161375 (NILs), non-climacteric parental lines.

(Perin et al., 2002) (Moreno et al., 2008)

Fruit flesh firmness Five QTL for flesh firmness in the cross Piel de Sapo x PI 161375 (NILs).

(Moreno et al., 2008)

Fruit flesh arroma profile Ester 3-hydroxy-2,4,4-trimethyl-pentyl 2-methylpropanoate: Two QTL in the cross Piel de Sapo x PI 161375 (NILs). (Z,Z)-3,6 nonadiena, responsible for the cucumber-like aroma: One QTL in the cross Piel de Sapo x PI 161375 (NILs). Octanal: One QTL in the cross Piel de Sapo x PI 161375 (NILs).


Root growth and architecture Seventeen QTL for root traits in the cross Piel de Sapo x PI 161375 (NILs).

(Fita et al., 2008)

Earliness. Nine QTL described in the cross Piel de Sapo x PI 161375 (F2 and DHLs). Three QTL for early fruit maturity in the cross Chinese line Q 3-2-2 x Top Mark (F2-F3).

(Monforte et al., 2004) (Cuevas et al., 2009)

Yield-related traits Four QTL for primary branch number, five QTL for fruit number per plant, four QTL for fruit weight per plant, two QTL for average weight per fruit and one QTL for percentage of mature fruit per plot in the cross USDA 846-1 x Top Mark (RILs).

(Zalapa et al., 2007)

Postharvest life traits Three QTL involved in reduced postharvest losses and 11 with a detrimental effect on fruits after storage in the cross Piel de Sapo x PI 161375 (NILs).

(Fernandez-Trujillo et al., 2007)

Sensory traits Thirty-two QTL including global appreciation, sweetness, sourness in the cross Piel de Sapo x PI 161375 (NILs).



Acknowledgements: The author would like tothank Dr. James McCreight for his critical review of thisgene list.

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